Webb Et Al 2002 Phylogenies and Community Ecology

Phylogenies and Community Ecology Author(s): Campbell O. Webb, David D. Ackerly, Mark A. McPeek and Michael J.
Donoghue Reviewed work(s): Source: Annual Review of Ecology and Systematics, Vol. 33 (2002), pp. 475-505 Published by: Annual Reviews Stable URL: http://www.jstor.org/stable/3069271 . Accessed: 04/12/2012 07:34
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Annu. Rev. Ecol. Syst. 2002. 33:475-505 doi: 10.1146/annurev.ecolsys.33.010802.150448 Copyright? 2002 by AnnualReviews. All rightsreserved Firstpublishedonline as a Review in Advance on August 14, 2002
AND COMMUNITY PHYLOGENIES ECOLOGY

David D. Ackerly2, MarkA. McPeek3, Campbell0. Webb1, and MichaelJ.Donoghue1
New Haven, 1Department Ecology and EvolutionaryBiology, YaleUniversity, of Connecticut06511; email: campbell.webb@yale.edu, michael.donoghue@yale.edu 2Department Biological Sciences, StanfordUniversity,Stanford,California94305; of email: dackerly@stanford.edu 3Department Biology, DartmouthCollege, Hanover,New Hampshire03755; of email: mark.mcpeek@dartmouth.edu
conservatism, Key Words community assemblyandorganization, phylogenetic nichedifferentiation biogeography, speciesdiversity, * Abstract As betterphylogenetic becomeavailable manygroups for hypotheses of organisms, studiesin community of ecologycanbe informed knowledge theevoby to lutionary relationships amongcoexistingspecies.Wenotethreeprimary approaches integratingphylogenetic informationinto studies of community organization: 1. examining phylogenetic the structure community of 2. assemblages, exploringthe basis of community niche structure, 3. addinga community and context phylogenetic to studiesof traitevolution biogeography. recognizea commonpattern phyand We of in and logeneticconservatism ecologicalcharacter highlightthe challengesof using of to phylogenies partial lineages.Wealsoreviewphylogenetic approaches threeemerof relative abundance and distributions, gentproperties communities: speciesdiversity, sizes. Methodological advances phylogenetic in characconstruction, range supertree terreconstruction, modelsfor community null evolution,and assemblyandcharacter metricsof community structure underlie recentprogress these arthe in phylogenetic eas. Wehighlightthe potential community for ecologiststo benefitfromphylogenetic knowledgeandsuggestseveralavenuesfor futureresearch.
INTRODUCTION
The differencesamong species thatco-occur in an ecological communityare the result of modificationsto a common ancestorthatthe species all ultimatelyshare. As molecularand analyticalmethods make the elucidationof phylogenetic relationships easier and more reliable, ecologists have an invaluablenew dimension of informationavailable with which to make sense of these differences among species. However, despite recognition of the potential for using phylogenies in community ecology (Brooks & McLennan 1991, Losos 1996, Thompson et al. 2001), and increasing interest in the role of history in ecology (Ricklefs 1987, Ricklefs & Schluter 1993a), integrationof evolutionarybiology and community
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WEBB AL. ET ecology remainselusive. This is due partlyto the conceptualand methodological difficultiesof bridginggapsof temporal spatialscale andpartlyto poorcommuand nication:many ecologists are eitherunawareof the potentialbenefitsof knowing about the phylogenetic relationshipsin their communitiesor are deterredby the unfamiliarityof moleculartechniquesand phylogeneticmethods and the accompanyingterminology.Similarly,many systematistsare unawareof the fascinating ecological questionsthatcan be addressedusing the phylogenies they produceor the ways in which knowledge of communitycomposition might bear on studies of characterevolution, diversificationrate, and historicalbiogeography.Our intention in this review is to introduceto both partiesthe various approachesthat have alreadybeen takento incorporate phylogeneticinformationinto community ecology. Phylogenies are being used extensively in the largerfield of evolutionaryecology (see Miles & Dunham 1993, Miller & Wenzel 1995, Ackerly et al. 2000), so we limit our review to studies and concepts explicitly relatingto the phylogenetic of andtaxonomicstructure local communities.We do not explicitlyreviewcharacter displacementin species pairs (Schluter2000a), adaptiveradiationin particular coevolution(host-parasite, clades (Schluter2000b), "host-client" plant-herbivore, or andhost-pathogen), generalhistoricalbiogeography, the uses of microbialphylogenies. Previousreviews and discussions of the interactionof phylogeny with et communityecology include Wanntorp al. (1990), Brooks & McLennan(1991, 2002), Eggleton & Vane-Wright (1994), Losos (1996), McPeek & Miller (1996), Grandcolas(1998), and Nel et al. (1998). Empirically,phylogenies and communityecology have been put togetherpreand dominantlyin studiesof communityassembly,organization, species co-occurthree majorapproaches(Figure 1). Other rence, and we identify in this literature questions of communityecology, relating to relative abundance,range size distributions,and species richness have received less attentionfrom a phylogenetic perspective,but we cover the work thathas been done so far. We then review recent methodologicaladvancesand conclude with suggesteddirectionsfor further work.
COMMUNITY STRUCTURE AND COEXISTENCE

Even thoughphylogenetic methods were developed fairly recently,a connection between taxonomyand communityecology has long been recognized: As species of the same genus have usually, though by no means invariably, the some similarityin habitsandconstitution,andalwaysin structure, struggle will generallybe more severe between species of the same genus, when they come into competitionwith each other,thanbetweenspecies of distinctgenera (Darwin 1859). Darwin's statementalreadycontains what we see to be the essential elements of that of an evolutionaryunderstanding communityorganization: species interact
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Character Evolution
Biogeography
Regional Local
TRAITS
Community Niche Structure
SPECIES ASSEMBLAGES
of to of Figure 1 Schematic summary variousapproaches theintegration phylogeny, the of astraits,andcommunities. Examining phylogeneticstructure community (1) semblages;(2) exploringthe phylogeneticbasis of communityniche structure; (3) contextto studiesof traitevolutionandbiogeography. addingcommunity
in communities,that species interactbased on their phenotypic differences and similarities,and thatphenotypicvariationhas a basis in evolutionaryhistory.In a of of syntheticunderstanding the originandmaintenance communitycomposition, threeelements are drawntogether:phylogeny,communitycomposition,and trait information(Figure 1). Researchershave tendedto approachthis synthesis using one (or more) of three methods: 1. analyzingcommunitytaxonomic or phylogenetic structure, exploringthe phylogeneticbasis of niche differentiation, 3. 2. and adding communitycontext to characterevolution and biogeography.We discuss these approachesbelow, in orderof increasinginformationrequirements inand creasingpotentialto reveal both ecology and evolutionin the past and present.
The PhylogeneticStructureof CommunityAssemblages

Key question: Is the distributionof species among habitats (or samples) in a communitynonrandomwith respect to phylogeny? (Using: species list of local community+ distributionof species among community samples + phylogeny of community species list) Since the adventof formalclassification,natural historianshave askedwhy differentareasare dominatedby differentspecies, genera,and families (e.g., Gentry
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WEBBETAL. taxonomic structureof communitieswas first addressed 1982). The quantitative Elton (1946), who reasoned that the lower numberof species per genus obby served in local areas than in the whole of Britain was evidence for competitive exclusion of ecologically similarcongenersin local habitats.Interestcontinuedin species/genus ratios for a numberof years (Moreau 1948, Williams 1964, Simberloff 1970, Tokeshi 1991) and was notable as the context for the first use of null models in ecology (Gotelli & Graves 1996). Implicit in these analyses was the same three-part interactiondiscussed above (Figure 1): communityorganization (i.e., the role of competition)can be deducedfrom the (assumed)ecological similaritywithin a genus, and the taxonomic structureof a community(i.e., the of significantdeparture species/genusratiosin communitysamplesrelativeto a regional species pool). Morerecently,the global consistencyof taxonomicstructure in forest communitieshas been examinedby Enquistet al. (2002), who compared the species/genusand species/familyratiosacrossmany standardized ha plots. 0.1 betweennumbersof generaor familiesand They foundan exponentialrelationship thenumbersof species, acrosstwo ordersof magnitudeof species number, sugand gest thatthisresultindicatestheexistenceof forces actingto constrain phylogenetic structure. The availabilityof phylogenies, along with methods for the constructionof and supertrees for assemblingthe phylogeniesof communities,now permitscomto can munitystructure be assessed phylogenetically.A simple logical framework then be employedto infer mechanismsof contemporary coexistence (Table 1, and see Figure2 for terminology).A clumpedphylogeneticdistribution taxa ("phyof indicatesthathabitat-useis a conservedtraitwithinthe pool logenetic attraction") of species in the community,and that phenotypic attractiondominates over repulsion. However,phylogeneticoverdispersion (repulsion)can resulteitherwhen closely related taxa with the most similar niche-use are being locally excluded repulsed),such thatthereis minimumniche overlapof coexisting (phenotypically species, or when distantlyrelatedtaxa have convergedon similarniche-use and Note thatthe fourthpossible interaction,phenotypic are phenotypicallyattracted. repulsionof traitsthat are convergent,will not tend to recreatephylogenetically clusteredcommunities,but phylogeneticallyrandomones. For example,Webb(2000) found thatthe tree taxa thatco-occurredin 0.16 ha Borneowere moreclosely relatedthanexpectedfroma random plots in Indonesian samplingof the local species pool. Assuming thatconservatismdominatesin the he this as evidence phylogenetic distributionof ecological character, interpreted for the predominant of habitatfiltering(andphenotypicattraction), opposed role as to local competitiveexclusion (and phenotypicrepulsion)of similarspecies. In a similar study,H. Steers (personalcommunication)determinedthat a measureof the frequencyof co-occurrence treespecies pairsin a Mexicandrytropicalforest of with theirphylogeneticproximity,againinterpreting this was positivelycorrelated as evidence of habitatselection for ecologically similar,phylogeneticallyrelated were species. Kelly (1999) found thatBritishplant taxa in extremeenvironments moreclosely relatedthanexpectedby chance,whichwas seen as evidencethatthese
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of taxa of distribution sample on thephylogeny a poolat a TABLE1 Theexpected and trait of combinations phylogenetic distribution scale,givenvarious spatial larger ecological process traitsphylogenetically Ecological Conserved Convergent force: Dominant ecological Clustered Habitat attraction) Overdispersed filtering (phenotypic exclusion repulsion) Overdispersed Random Competitive (phenotypic
species were ecologically similar.Conversely,Graves & Gotelli (1993) showed that congeners seldom co-occur in the same mixed-species foraging flock in the Amazon, but that this "checkerboard" patternbreaksdown at higher taxonomic this levels. They interpreted finding as the effect of intra-community competitive exclusion among ecologically similar species (i.e., phenotypic repulsion), with congeners being most similar.In Florida woodland communities dominatedby oaks, J. Cavender-Bares (personalcommunication)also found thatclose relatives co-occurredless thanexpectedby chance.In this case each plot generallyhad one species from each of threemajorQuercusclades (sections). The spatialscale of samples used in studies of communityphylogenetic strucof tureis of great importanceto the interpretation the patternsfound because the and biological natureof phenotypicandphylogeneticattraction repulsiondepends uponthe scale involved.At the largest,continentalscales (e.g., 1,000-10,000 km), phylogenetic clusteringof membersof a regional sample on a global phylogeny reflectsbiogeographicratherthanecological processes, as clades diversify within the sampleregion,andcause manytaxain theregionto be, on average,morerelated to each otherthanto taxa outside the region. Withina region (e.g., 10-1,000 km), phenotypic sorting might occur among communitiesthat differ environmentally from one another (e.g., wetlands versus montane). Such phenotypic attraction or might lead to phylogeneticattraction repulsionof the communitysampleon the traits.Susof regionalpool, dependingon the phylogeneticdistribution important tainedphenotypicrepulsionwithina communitymightalso lead to semipermanent exclusion of too-similartaxa from individualcommunities,with taxa maintained in the regional pool by low rates of dispersalamong communities(e.g., Tilman 1994). At the community scale (e.g., 100 m-10 km), species should segregate into habitatsbased on the relative strengthsof habitatfiltering versus competition among similar species (see Figure 2). Finally, at the smallest, neighborhood interacscales (e.g., <100 m), one might observe the effect of individual-based or "neighborhood exclusion." Hence, a tions that lead to within-habitatfiltering may detectdifferent spatiallynestedanalysisof communityphylogeneticstructure of phylogenetic clusteringor over-dispersionat differentscales, providpatterns ing more informationaboutcommunityprocesses thanan analysis at just a single scale.
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intrinsictrait:x
PHYLOGENY
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1 x BIOGEOGRAPHY
2 y
3 x
4 y
5 z
6 z
7 z
Species Traits
1 RegionI
3 (i, ii) i
Species e Region/I
(iv) Community Assembly:

-
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Fundamental Niche Y Fundamental NicheZ ^ ECOLOGY
E[4I2
(5
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(iii) RealizedHabitat: Y Zi 5 z Z1 Z1 6 z Z1 Z2 7 z Z2
Species COMMUNITY Traits PHYLOGENETIC 'Intrinsic' STRUCTURE 'Extrinsic'Traits (e.g. observed habitat)
2 y Y
Figure 2 Schematic of the general framework employed in this review, with associated terms. A lineage may diversify by the division of its ancestral range and allopatric speciation, such that sister clades are no longer coregional (ancestor of species 1 and 2 versus ancestor of species 3-7; BIOGEOGRAPHY).Alternatively, sympatric and parapatricor even allopatric speciation mechanisms may lead to the origination of new species that are coregional with their sister species (species 3-7). A phylogeny can be reconstructedfor the lineage (PHYLOGENY)using molecular and morphological species traits. Species may appearin regions either through the geographical division of their area (vicariance) or by subsequentdispersal (species 2
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into dispersed regionII). The phylogeniesof lineagesand Figure 2 (Continued) of the distribution taxa amongregionscan be used to inferthe historicalpatterns in of movement the taxaandassociated reconstruction areaconnectedness of (area methods.Traitchangeoccursas the lineagedicladograms), usingbiogeographic state on and versifies, ancestral changescanbe reconstructed traced) a phylogeny (or or likelihood(x to y, y to z). Traitscan using, for example,parsimony maximum or traitsthatcanbe usefullybe dividedinto intrinsic (morphological physiological is assessedvalidlywhen an organism removedfromits environment, e.g., skeletal structure, beak size, body size, plant sexual system) and extrinsic(traitsthat only have meaningin an externalabioticandbiotic environment, swimming e.g., shadetolerance, tolerance, speed,maximum growthrate,drought preychoice,diet of The "ecologicalcharacter" an organismis usuallya complex set of breadth). but correlated characters, canoftenbe directlyrelatedto simplerintrinsic morphologicalcharacters (e.g., Losos 1995). The pattern evolutionof any traitcan be characterized conservative of as (more the closely relatedtaxa are more similar)or convergent (homoplasious, indepenTraitdivergence not intrinsically dent evolutionof similarity). is conservative or but becausethe greaterthe changein a trait,the more likely it is to convergent, resemblethe value of a species in an independent lineage, divergenceoften resultsin convergence. therateof homoplasy increases withdecreasing Additionally, number potential of traitstates(Donoghue Ree 2000) andincreasing & of number for speciesto be functionally similar whileintrinsically different ways (e.g., fleshyfruitedness of arisesvia a number anatomical paths).Whenreferingto ecological nicheconservatism" beenused (Harvey Pagel has & traits,the term"phylogenetic Lordet al. 1995).This conservatism be due to active,stabilizing selec1991, may tion(Lord al. 1995)orto a reduction thepotential et in of ecologicalcharacter range evolutioncausedby the fixationof ancestral traits(i.e., developmental constraints; Westobyet al. 1995). from a regionalspecies pool (the list Species are assembledinto communities of all species in an area at the next-highestspatialscale from the scale under consideration; e.g., species 2-7 form a species pool relativeto the community) Liebold 1998, Fox 1999, Blackburn Gaston2001) (ECOLOGY). this & In (e.g., reviewwe considera community be morethantwo speciesin the sametrophic to level and the same guild (e.g. meadowplants, desert granivores) co-occurring spatiallyat a scale over which species might dispersewithina few generations. Some species presentin a region may fail to meet the ecological requirements to survivein any of the niches in a community, is, they are filteredout, or that fail to reach a particular chance (species 3 and processi simply community by or ii, respectively). with regional Community species richnessis often correlated pool richness,suggestingthat communitiesseldom saturate(Cornell& Lawton 1992). Species may persist (coexist for long periods) in the same community by occupyingdifferentniches, thus minimizingcompetitionfor resources
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ET WEBB AL. ("habitat Figure2 (Continued) partitioning"; species2 and4 versusspecies5-7; with several Wisheu1998).However, enoughspeciesin thecommunity, specieswith nicherequirements tendto be filtered the sameniche(a phenotypic will into similar interaction attraction). negativeinterindividual (phenotypic repulSimultaneously, habitat is expectedamongspeciesthatoccupythe samenicheand/or sion) (process is iv). This negativeinteraction expectedto be stronger amongecologicallymore of distribution taxaintosubsimilarspeciesandmayaltertherealized niche/habitat niche (butneverbeyondthe bounds niches (or localities)withinthe fundamental of the fundamental niche;species 7 versus5 and6), or may lead to classic comexclusion(processiii on species 4). It may also lead to the exclusionof petitive scale (e.g., at the scale neighborhood competingspecies on the most proximate, withoutleadingto community exclusion.Speciesmay of individual interactions), coexistwithina habitat non-niche-partitioning, (sensuChesson2000) by equalizing andprocessv), ormaybe in theslow,nonequilibrial process processes(species5, 6, We to of beingexcludedfromthecommunity. use habitat meanthe spatiallocation of levels come together. wherea combination resource in for definedarea A phylogenycanbe reconstructed speciessampled a spatially PHYthanfor all speciesin a lineage(COMMUNITY rather (region,community), The occurrence taxain areasat a smallerspatial of LOGENETIC STRUCTURE). on and can scale (community habitat, respectively) be indicated the phylogenyof of the largerpool of species. The distribution these taxa can be phylogenetically on clumped,random,or over-dispersed the phylogenyof the entirepool (e.g., a on 6 and7 is clustered the phylogenyof the community species samplecontaining of species2, 5, 6, and7). Sustained selective pressure from individualcompetitorsor environmental over to time, changesleadseventually changein ecologicalcharacter evolutionary The in the direction reducednicheoverlap(EVOLUTION). community of usually contextmayalso causestabilizing selection,especiallyin diversesystemsin which Traitchangemay also be the resultof are interactions unpredictable. interspecific Traitchangeresultingfromecologicalinterdrift,foundereffects, or pleiotropy. actions may itself lead to ecological speciationwithin a region (McPeek 1996, Schluter 2001).
The PhylogeneticBasis of CommunityNiche Structure

arrayedon a phylogeny? Keyquestion:How are niche differencesin communities list of local community distribution species withincommunity + of (Using: species + phylogeny of communityspecies list + ecological character data for those species) of of Ecologists have long studiedthe distribution ecological characters species MacArthur Levins & in communitiesto understand (e.g., communityorganization 1967, Bowers & Brown 1982). An even dispersion of trait values along some
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gradient(e.g., constantbody-size ratios) has been held to be evidence for both contemporarycompetitive exclusion and the long-term evolutionary effect of such competition(Connell 1980), althoughthe establishmentof the significance of this overdispersionhas been controversial(reviewed by Gotelli & Graves 1996). of The availabilityof dataon the taxonomicor phylogeneticdistribution niche niche structureto either condifferences enables the allocation of contemporary temporaryecological or historic evolutionarycauses, or a combinationthereof. For example, because Cavender-Bares (see above, personalcommunication)had in assessedwater-usetraitdataas well as wateravailability sampleplots (Cavenderthe Bares& Holbrook2001), she was able to interpret phylogeneticoverdispersion habitatfilteringmediatedby characters of local plots as the resultof contemporary that were phylogeneticallyconvergent.Similarly,Webb & Peart(2000) assessed the species, genus, and family associations of rain forest trees with three habitat types and found both genera and families that had all of their species associated with a particularhabitat.Although not phylogenetic, this patternof ecological of conservatismsupportsWebb's (2000) interpretation the role of habitatfiltering. In the same forests, Ashton (1988) has arguedthat too much emphasis has been placed on differences in habitatuse among species and that in the Diptewith rocarpaceae, genera and sections differ significantlyin ecological character, floral) species within them differingmainly by minormorphological(particularly changes. At a deeper phylogenetic level, Dimichele & Phillips (1996) showed fossil plant comconsistent occupationof varioushabitatsin Pennsylvanian-age munitiesby differentlineages (lycopsids, seed plants,ferns,andsphenopsids),and T. Feild and colleagues (personalcommunication)have inferredthe maintenance in of an ecological niche (disturbed understory) severalearly angiospermlineages. Whereasclear cases of simple conservatismoccur,it is likely thatwith a large of communitythephylogeneticdistribution traitsis a complexmix of conservatism and convergence.A differentphylogeneticscope of a study (e.g., a communityof Quercusversus all angiosperms)might thus have a stronginfluence on the comobserved.Silvertownet al. (1999) demonstrated for munityphylogeneticstructure meadow plants that mean pairwise co-occurrenceof species in a hydrologically defined niche-space was less than expected, indicatingsignificantdivergencesin habitatuse among species. Using the same methods, comparingjust the species within a genus, they found a few cases of segregation of species, some cases with randomoverlap, and some examples of higher than expected niche overlap (Silvertownet al. 2001). This mixtureof overlapand segregationwas also seen at higher taxonomic levels, althoughthe broadestcomparisonshowed eudicots and monocots to be more segregatedthanexpected.These resultsindicatethatthe underlyingecological traitsexhibitedvaryingpatternsof divergenceand stasis (i.e., convergenceand conservatism)correspondingto the observationof segregation and overlap, respectively. In a related study of community assembly, Tofts & Silvertown (2000) assessed the effect of environmentalfiltering on grassland communitymembership,findingthattraitvalues for species in a local community
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were less variable than those in a regional pool, independentof phylogenetic contraststo removethe effect of phyeffects; they used phylogeneticindependent logenetic traitconservatism. The trait-phylogeny-community relationshipmay also appearto be random: Winston(1995) foundno differencein co-occurrenceratebetween groupsof phylogenetically closely related and more distantlyrelated streamfish, even though co-occurrencewas less among morphologicallysimilar species than among less to similarones, a resulthe attributed the effects of competition.Barraclough al. et (1999) tested whether species of tiger beetle that co-occurredin the same habitat at a locality had lower than expected similarityin various ecomorphological characters,using a phylogeneticallybased null model. They found no evidence for character divergencebetweenco-regionalor co-occurringspecies, or for habitat divergencein co-regionalspecies. Divergencepatternswere indistinguishable from randomcharacter change (see also McCallumet al. 2001). Some studiesperformthe equivalentof phylogeneticindependent contrastanaltraitsandcommunity ysis (Felsenstein1985) on the associationbetweenparticular For membership. example,Chazdonet al. (2002) askedwhethertherewere associtraitsandforesttypes for CostaRicantrees,andfound ationsbetweenreproductive thatthe significantassociationsfound could be explainedby the differentphylogenetic composition of the forest types and the generally phylogeneticallycontraits.Ibarra-Manriquez al. (2001) showed servativenatureof the reproductive et the same outcome for differences in seedling germinationtype among different of forests. Otherstudies have looked primarilyat the distribution ecologically related traitsof species from a species pool, withoutreferenceto community-level variationin species composition (Grandcolas1993, Brandlet al. 1994). For ex(1999) found little evidence of conservatism ample, Bohning-Gaese& Oberrath in ecological traitsfor 151 co-occurringbird species.
BiogeographicHistory,CharacterEvolution, and CommunityAssembly

conditionsdidpatternsof nicheKeyquestion:Whereand underwhat community local communities+ distributionof species use originate? [Using: species list of within communities+ phylogeny offull lineage (and of other lineages) + ecological trait datafor lineage members+ biogeographicreconstructionof regional distributionof ancestral taxa] When phylogenetic and ecological informationis availablefor all extanttaxa in a lineage, includingspecies in and outside of a particular community,ancestral of character reconstruction ecological traitsandniche use can be examined(within et limits of reconstruction methods;Cunningham al. 1998). The questionsof nicheuse evolutiondescribedabovecan thusbe answeredmoreaccurately(e.g., McPeek & Brown 2000). Including all taxa in a lineage permits assessment of the contemporarygeographic distributionof clades. Where all the members of a clade are currently
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co-regional,and no membersof other (closely related)clades are present,the reand evolution gion maybe consideredto be closed (withoutimmigration), character the Furmethodscan also be used to reconstruct evolutionof communitystructure. interactionsoccurred where there is evidence that (a) intra-community thermore, among all taxa in the region (often assumed) and (b) no majorinteractorsin the historical communities came from other clades (e.g., ants versus rodents), trait as changes can be interpreted both cause and effect: The ancestralcommunities can as can be reconstructed changein characters be interpreted the responseto and characterstates in other ancestralspecies. Workby Losos and colleagues (Losos AntilleanAnolis lizardsexemplifiesthis 1992, 1995; Losos et al. 1998) on Greater the majorityof species on each islandresultfromintra-island radiations, approach: and a similarpatternof habitatuse (e.g., crown, trunk,twig) has evolved on each island within each radiation.This method also allowed the researchersto reconin structthe compositionof ancestralcommunitiesandto inferthatdiversification habitatuse was a responseto the habitatsalreadyoccupied. Most regions are, however,open (Losos 1996), i.e., they contain membersof clades thatcan be inferred(using biogeographicmethods)to have originatedelsewhereandto have arrivedin a regionby dispersal.Forexample,McPeek& Brown the (2000) reconstructed phylogenyof all Enallagma(damselfly)species in eastern North America and were able to differentiatebetween a recent radiationin New England and an older diversificationin the southeastUnited States; most eastern lakes from the Gulf of Mexico to Canada,however,containmembersof both clades. Combiningsuch biogeographicdatawith traitreconstructions permitsthe of determination whethertraitsandcurrent niche-useevolved in situor whetherthe traitswere establishedelsewhere,outsidethe current region(Brown& Zeng 1989). of Cadle& Greene(1993) analyzedthe clade compositionand size distribution 15 in Neotropicalcolubridsnakecommunitiesto test whetherthe overalldistribution sizes in any communitywas independentof the clade composition. They found that (a) the overall size distributionof a communitybore a direct relationshipto the clade composition, (b) different clades had different mean sizes (and sizeassociateddifferencesin diet), and (c) the lineages appearto have originatedand diversifiedin isolated areas.The workthus indicatedthe role of historyin shaping the regionalpool of species anddid not supportthe hypothesisthatsnakecommunities are organizedby processes that tend to maintainsome overall distribution in size. In his study of Lesser AntilleanAnolis species, Losos (1992) noted that the one or two species on each island are not generalists,as the ancestorsof the Greater Antilles lizardsare inferredto be, but show conservatismof niche-use and appearto have been ecologically sortedonto the islands accordingto the available habitats(see also Vitt & Zani 1996, Zimmerman Simberloff 1996, Sturmbauer & 1998, Barker& Mayhill 1999, Craiget al. 2001, Galassi 2001). Richman(1996) used phylogenies to suggest that Japanese and Europeanassemblages of Phylloscopus warblerswere formed by multiple invasions from mainlandAsia, with only limited subsequentspeciationand little associatedmorphologicaldiversificain tion. Therewas strongconservatism body size throughout genus, irrespective the
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of region, and no evidence of convergence in morphology or habitat-use in Europe and Japan.In addition (in contrastto previous analyses, e.g., Richman & Price 1992), habitat-use(low-elevationconifer versus high-elevationdeciduof ous) divergencewas shown to have mainly occurredearly in the diversification the lineage (althoughhabitat-use only inferredfrommorphology, recorded was not directly). It is often notedthatthe ecological character species in today'splantcommuof nities reflects the biogeographichistory of the species and theirrecent ancestors. For example, Lechowicz (1984) asked why temperatetree communitiesshow a wide range in the time of leaf appearancein spring,using physiological, phylogenetic, and biogeographicdata.Therewas no significantphylogeneticclustering of early versus late leafers (based on taxonomiesavailableat the time), but early leafers were generallyderivedfrom cool temperatelineages, whereaslate leafers were primarilyof tropicallineage origin, reflectingevolutionaryconservatismin montaneforests of Mexico the everphysiological traits.Similarly,in subtropical is greenunderstory composed of tropicalelements, while the deciduousoverstory is composed primarilyof species from temperateclades (Williams-Linera1997; see also ter Steege & Hammond2001). ecosystems have been the subject of considerablestudy Mediterranean-type owing to the apparentconvergences in plant and animal communities on five continents (Cody & Mooney 1978). Community-levelconvergencein distributions of ecological traitsmay arise by a combinationof recentadaptiveresponses of the organismsto currentenvironmentsand by sortingor assembly of lineages drawnfromthe regionalspecies pools. Phylogenetic,biogeographic,andhistorical In studies are key to evaluatingthese alternatives. the plants of Andalusia,Spain, Herrera that (1992) demonstrated character syndromesdifferedbetween"old"and "new"lineages (distinguishedby fossil recordsandintercontinental disjunctions). The persistenceof ecological characterin old lineages again emphasizesthe imto Expanded portanceof conservatismin traitscontributing communitystructure. and betterresolvedphylogeniesnow makeit possible to evaluatesuch hypotheses with detailedstudiesof ancestralstatesin relationto communityassembly.In California chaparral, D.D. Ackerly (unpublished)found thatplants with similarleaf in flora were derivedfrom both subtropicaland characteristics the contemporary ancestral north-temperate lineages.In this case the subtropical lineagesmaintained traitsthat predatedthe mediterranean-type climate, while the temperatelineages to exhibitedmorerecentevolutionaryshifts, presumably adaptations representing climatic conditionscomparedwith theirbiogeographicorigins (see also changing Verduet al. 2002). Reconstructingancestral communities and determining the causes of trait change is harderin open regions than in closed ones, but methods have been developed for this. When several lineages share the same area cladogram and can be assumed to have evolved in the same time frame, co-regionalitycan be inferredfor ancestralspecies (co-speciation sensu Brooks & McLennan 1993). If community co-occurrenceand biotic interactioncan be reasonably assumed
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between membersof differentlineages, patternsof traitchangecan be interpreted as co-adaptations(sensu Brooks & McLennan 1993). Co-adaptation may most be inferredwhere traitchange in one lineage is associated with putareasonably tive dispersalinto a region by a species in anotherlineage (see figure 3 in Losos the 1996). Brooks & McLennan(1991) reconstructed influenceof pre-adaptation andco-adaptation helminthparasitecommunitiesin Neotropicalstingrays,conin sidering differenthost species as differentniches at six different sites in South America.They showed thatthreecontemporary communitiescame aboutprimarvicariant division of ancestralcommunities, two originatedby dispersal ily by of parasites into a region without switching host species (or niches), whereas one representeda complicatedassembly of vicarianceand dispersal,with some host conservatismand some host switching (see also Poulin 1999, Sasal et al. 1999, Morand& Guegan 2000). Mayden (1987) used geological and fish phyto the logeny information reconstruct historyof vicariancein Ozarkriverdrainages. Gorman(1992) then added informationabout habitatuse (position in water colancestral umn)of contemporary species to reconstruct communitycompositionand habitatoccupancyandwas able to infer which competitiveinteractions were leadniche displacement,which he tested experimentally.Losos ing to contemporary et al. (1997) also appliedanexperimental to approach test hypothesesderivedfrom biogeographicand trait evolution data, finding that Anolis limb length evolved adaptivelyin populationsestablishedon small islands.
Community Organization:Conclusions
A phylogeneticapproach studyingcommunityorganization to providesa new perspectiveon the perennialquestionsof the role of competitionandthe maintenance of diversityin communities,by highlightingthe similaritiesof co-occurringspecies as well as the differences.A dominant perceptionin evolutionary ecology is thatcoexisting species must differ significantlyand that most variationbetween closely related species is the adaptiveresponse to past competitionwhen species did not differ (e.g., Harvey & Rambaut2000). The general predictionfrom this model of evolutionis that divergence(and thereforehomoplasy) in ecological character should be widespread.However,phylogenetic analysis reveals that many (possibly the majorityof) lineages studiedshow evidence for conservatismof dominant ecological character,in both animal lineages (e.g., Richman 1996, Barraclough et al. 1999, Petersonet al. 1999, Lindeman2000, McPeek & Brown 2000, Price et al. 2000, Forstmeier al. 2001) and in plants(Grime 1984, Peat & Fitter 1994, et Lordet al. 1995, Ackerly 1999, Prinzinget al. 2001). For plantsthis runscounter to a long-standingbelief that functionaland ecological strategiesevolve rapidly, between taxonomic leading to widespreadconvergenceand little correspondence and ecological groupings(e.g., Warming1909, Cronquist1988). An associatedinsightarisingfroma phylogeneticperspectiveis thateven if convergentevolutionhas occurredin a single trait(e.g., diet), otheraxes of ecological similarity(e.g., forest-typeuse) are often conserved.The moretraitsinvolved,the
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WEBBETAL. more likely it is that a composite measureof "net ecological similarity"will be conservedin a lineage, especially if thereare life-historytrade-offsamong traits. Change in such a composite measurewould perhapsbe best modeled by evolutionarydrift. Ecologically, difference on one niche axis alone may be sufficient to reduce competitionwith other species under stable conditions, but it is more shifts overtime, with species experiencing likely thatthe competitiveenvironment interactionswith other species on differentniche axes (food, shelter, intermittant water,nutrients,space). Hence, when viewed across many traits,interactionsthat challenge the co-existence of closely related,ecologically similarspecies may be more frequentthanrecognized. If conservatismof ecological traitsis widespread,then the effect of contemporarycompetitioncan be assessed by the extent to which phylogeneticallyrelated, ecologically similarspecies co-occur:avoidenceof closely relatedspecies by one another(e.g., Graves& Gotelli 1993) would be indicativeof strongcompetition among similarspecies. Because the strengthof negativeinteractionshouldgenerto ally be proportional the phylogeneticproximity,a likely communityoutcome patternof bothphylogeneticclumpingandoverdispersion: mightbe a hierarchical Some conserved characterswill determinethe ability of taxa to occupy a fundaof mentalniche (leading possibly to overall phylogeneticattraction co-occurring taxa,e.g., taxa 5, 6, and7 in Figure2), whereasotherswill cause local competitive exclusion (leading to phylogenetic repulsion of co-occurringtaxa within those attracted clades, e.g., taxa 6 and 7 in Figure 2). Whererelated,similarspecies do co-occur, attentionmust be given to mechanismsthat permitthe co-existence of similarratherthandifferentorganisms(e.g., Chesson & Warner1981, McPeek & Brown 2000, Hubbell2001). on The studyof adaptiveradiations islands (e.g., Givnish 1998) may have led to character on an overemphasis evolutionary displacementandecological niche partitioning.In diverse, continentalcommunities(in which most species live), interover time, both on the timescaleof individuals actionsmay be moreunpredictable (becauseof the diversityof neighbors)(Connell 1980), andbecausespecies ranges may change more often. This would lead to weakerpairwise interactionsamong sistertaxa, and species radiationsmight occurwith little ecological differentiation et (sensu McPeek & Brown2000, e.g., Richardson al. (changebeing nonadaptive) 2001). Local communitieson continentsmay then representa selective sampling of the regionalpool to minimizeecological similarityor may only be able to contain species that are similar (over long periods of time, if not indefinitely). On islands, species must change or go extinct. A systematicreview of the prevalence of conservatismand convergencein both island andcontinentalsystems would be very valuable. new perspectivesbut phylogeneticinformationoffers important Incorporating also brings additionalchallenges. Perhaps the most immediate challenge is to explicitly define the ecological, spatial,and taxonomic scales in a study because the processesthatstructure assemblyof regions,communities,andhabitatsdifthe of fer (see ThePhylogeneticStructure Community Assemblages,above),andusing
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phylogenies only of taxa that co-occur at a particularspatial scale can confound inferencesaboutecological andevolutionary processes (see Jablonski& Sepkoski it is valid to assess the correlationbetweenrelatednessand simi1996). Although larity (i.e., the effective degree of conservatism)in any sample of species, it must be rememberedthat partial(community)phylogenies may not provide accurate evolution or lineage-wide assessmentof conservatism inferences aboutcharacter and homoplasy (the issue of "taxon sampling";Ackerly 2000). An example is given in Figure 2, in which the taxa includedin the communityphylogeny would of lead to an incorrectreconstruction the evolutionof traitsy and z. We recognize that all phylogenies of extant taxa are only partialsamples, owing to extinction, but phylogenies for co-occurringspecies are particularly egregious samples and are likely to introducesystematicbias in the study of traitevolution owing to the of in distribution ecological characters differentcommunities. estimates of the absolute lengths of branchesin a partial(commuObtaining nity) phylogeny greatly increases the accuracy of inference about the correlation betweenrelatednessand similarity.Estimatesof relativebranchlengths often come fromratesof molecularevolutionbutultimatelydependuponthe datingand phylogenetic placement of fossils. Age estimates are also vital for the accurate of reconstruction the species compositionof ancestralcommunities.Congruence of the topologies of differentlineages without age informationcannot determine the order of arrivalof taxa in an ancestralregion or the temporalorder of trait change in the lineages. Disentanglingcause from effect requirestemporalinformation (see Donoghue et al. 2001, Hunn & Upchurch 2001, Sanmartinet al. 2001). A relatedchallengewhenusing regional-scalebiogeographicmethodsto reconinteractions thatcommunityco-occurrenceof structhistoricalintra-community is ancestraltaxamustbe assumed.The spatialscale of the areasin an areacladogram is usually far largerthan a single community(Grandcolas1998), and coregional ancestralspecies may never have interacted,because they occurredonly in differenttypes of communitiesor because they were spatiallysegregatedby chance while still sharinga region.However,phylogeographic methods(Avise 2000) may to historicalpopulation permitthe use of intra-speciesgenetic variation reconstruct historicalco-occurrence(e.g., movement and to indicate patternsof intra-region Zink 1996, Comes & Kadereit 1998, Taberletet al. 1998). Fossil communities (e.g., Wing et al. 1993, Hadly & Maurer2001) might also be used to confirm communityco-occurrence.
EMERGENT PROPERTIES COMMUNITIES OF

Most work to date at the intersectionof phylogenetics and communityecology has dealt with the presence and absence of species in a community,associated differencesin traits,and the assembly of communitiesthroughtime. Community ecology also deals with species diversity,the relative abundanceof co-occurring
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WEBB AL. ET of species, and the distribution range sizes. Therehave been a numberof creative uses of phylogeny in these areas.
SpeciesDiversity
Key question: Whydo differentareas vary in the species richness of particular clades? (Using: species lists of local communities+ local and global estimatesof species richnessof clades + phylogeneticrelationshipsof local and global clades) One answerto the question"Whydo similarhabitatsin differentregions have differentnumbersof species?"is thatdifferinghistoriesof the areashaveled to occupancyby differentclades (Latham& Ricklefs 1993, Schluter& Ricklefs 1993, Qian & Ricklefs 1999, Ricklefs 2002) and thatdifferentclades have differentpotentialsfor diversification (Farrellet al. 1991, Sanderson& Donoghue 1996, Dodd et al. 1999, Gardezi& da Silva 1999) andthusdifferentnumbersof extantspecies. A second answeris that the areasdiffer in the length of time they have been occupied (Brown et al. 2000, Ricklefs 2002). For example, a latitudinalgradientin diversitymay reflectthe relativeages of majorclimateregions (andthe challenges of adaptingto temperateclimates) as much as the effect of local ecological processes (Blondel & Vigne 1993, Latham& Ricklefs 1993). Phylogeneticanalyses and age inferences are centralto testing such hypotheses:For example, Ricklefs & Schluter(1993b) found thatthe clades of passerinebirds in Panamanian forest were on average2.6 times the age of the clades in forests in Illinois. The dominant hypothesisfor why islandsdifferin species richnessis MacArthur & Wilson's (1967) equilibriumtheory of island biogeography.Phylogenies are playing a majorrole in testing the applicabilityof this hypothesis.Using molecular estimates of species age, Ricklefs & Bermingham(2001) were able to reject the assumptionof constantrates of immigrationand extinction in Antillean birds. Using island species lists and a phylogeny,Losos & Schluter(2000) were able to differentiatebetween in situ speciationof Anolis lizardson Caribbeanisfrom otherislands.They showedthaton islands largerthan lands,and immigrants 3,000 km2,in situ speciationovertakesimmigrationas a source.
RelativeAbundance
Keyquestion:How does the relativeabundanceof taxa vary across a community + phylogeny?(Using: species list of local community relativeabundancestructure + of local community phylogenyof species in local community) The distribution relativeabundance communitieshas long been the subject of in of attentionby ecologists. Whereasthere are many ways to statisticallycharaccurves (e.g., log-series, log-normal), terize the shape of abundance-distribution models have been few (but see MacArthur1960). Fewer satisfactoryexplanatory The still make predictionsaboutthe relationof phylogeny to relative abundance. most comprehensive model of communityabundance (Hubbell2001) predictsthat and widespreadthan younger species (in older species should be more abundant itself, an old idea; Willis 1922). Methodsexist to date nodes in a phylogeny,but
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the pruningof phylogenies by extinction means the time since divergencefrom the most closely relatedextanttaxon may often be a poor predictorof species age (Box 2 in Chown & Gaston2000). An increasing number of studies have asked if there is a relation between characters a taxonomicgroupand the mean abundanceof its members(Farrell of et al. 1991; see also Heard& Hauser 1995, Edwards& Westoby2000, Murray& Westoby2000). Schwartz& Simberloff(2001) found thatvascularplant families with few species tended to have fewer than expected rare species. Such analyses will benefit from an explicit phylogenetic framework:Webb & Pitman (2002) foundthata rank-based associationbetweencommon species anddiversefamilies of rain forest trees disappeared when consideredphylogenetically. Anotherapproachto relativeabundanceis illustratedby the work on bird size and abundance(e.g., Cotgreave& Harvey 1994, Harvey& Nee 1994). In Britain, overall bird populationsize is negatively related to body size (Nee et al. 1991), but within a tribe the relationshipis often positive. If ecological similarity and competitionare correlated,and if largerbodiedbirdsattainhigherdensitiesunder competitionthansmallerbirds,thenthose cladesthatcontainthe most ecologically similar species should show the strongestpositive association of body size and abundance. Nee et al. (1991) foundthatthe branchlength(using taxonomiclevels) from the base of a tribe'sclade to the rest of the birdswas a good predictorof the strengthof the positive relationship,longer branchesbeing associatedwith more complete guilds. Examining relative abundancestructurefrom a phylogenetic standpointwill surely be an exciting avenue of research.Finding any associationbetween abundance and relatednesscould indicate that local abundanceis actively influenced by phylogeneticallyconservedcharacters.
GeographicalRange
Key question: How do the sizes and spatial arrangementof species ranges vary across a phylogeny? (Using: species lists of local communityor region + range information these species + phylogenyfor these species) for and Range size can be treatedas a continuouscharacter, its evolution and association with other charactersassessed. The communitycontext of the species in such analyses is generally not addressed.In the few cases studied,closely related species tend to have more similarrange sizes than distantlyrelated species (Jablonski 1987, Ricklefs & Latham 1992, Brown 1995). This might appearto imply that (a) some ecological traitsare responsiblefor range size and (b) there is some conservatismin these traits.However,an alternative explanationfor such a patternis that related species tend to be of a more similarage than less related species, and if age is correlatedwith area (Willis 1922, Fjeldsa & Lovett 1997), then areawould appearas a phylogeneticallyconservedattribute species. Kelly of & Woodward(1996) investigated the correlationbetween life-form and range size in British plants, using a phylogeny, and found that trees have largerranges than nontrees, and that wind-pollinatedspecies have larger ranges than related
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non-wind-pollinated species. Gregory (1995) found that phylogenetic conserbetweenrangesize andbody size in British vatismdid not explainthe relationship contrastsin an analbirds.Gotelli & Taylor(1999) used phylogeneticindependent the probabilityof stream colonization by fish with body size, ysis correlating populationsize, range size, and distance from range center.They found that the importanceof removing phylogenetic effects to detecting the effect of distancewas from-range-center substantial. Range informationhas also been used to investigatemodes of speciation,reasoning in the "oppositedirection"from most work reviewed here (Lynch 1989, et Barraclough& Nee 2001, Gimaret-Carpentier al. 2002). Species range size is likely to be closely linked to the probabilityof furtherdiversificationunderany model of allopatricspeciation (Barraclough& Vogler 2000, Chown & Gaston et 2000). Barraclough al. (1998) used the range overlapof clades as an indicator of the mode of speciationin tiger beetles. If allopatricspeciationand subsequent range movement dominated,the degree of range overlap between sister clades should start low for two sister species and increase as more taxa are included in both clades. However, sympatricspeciation and subsequentrange movement shouldstartwith high rangeoverlapof sisterspecies anddecreaseas moretaxaare included.The authorsfoundthatoverlapstartedlow andincreasedwith increasing clade inclusivenessand inferredallopatricspeciation.
ADVANCES METHODOLOGICAL Community Phylogenies

Phylogenetic methods are in a sustainedphase of rapid development,with new maximum-likelihood(Lewis 2001) and Bayesian (Huelsenbecket al. 2001) approaches being explored. Phylogenies from separatestudies can now be joined et to form "supertrees" (Sandersonet al. 1998, Bininda-Emonds al. 1999), either being assembled "by hand"(e.g., Donoghue et al. 1998) or using algorithmsto resolve conflict amongtrees (Semple & Steel 2000, Salaminet al. 2002, BinindaEmonds et al. 2002). From these supertrees,phylogenies of communityspecies lists can be prepared(e.g., Tofts & Silvertown2000, Webb 2000); tools are now availableto facilitatethis process (Webb& Donoghue 2002). Communityphylousually lack informationaboutbranchlength, genies constructedfrom supertrees but there are supertreemethods that yield branchlengths (Lapointe& Cucumel 1997), and we anticipatethatbranchlengths based on absoluteage estimateswill soon be availablefor many groups (e.g., Magallon& Sanderson2001).
Testsfor PhylogeneticConservatism
of As we have emphasized,predictionsandinterpretation patternsof phylogenetic structuredepend on patternsof ecological similarityand divergence community among related species. Using taxonomic information,these patternshave been variation evaluatedwith hierarchical interspecific analysisof variance,partitioning
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into differentlevels: species withingenera,generawithinfamilies, families within orders, etc. (e.g., Mazer & Wheelwright 1993, Peat & Fitter 1994, Lord et al. of 1995). Withthedevelopment well-resolvedphylogenies,a varietyof quantitative methodshave been proposedto examine the extent to which ecological traitsare conservedor convergenton a phylogeny.For discretecharacters, randomizationbased tests of the number of reconstructedcharacterchanges have been used (Maddison& Slatkin 1991, Barracloughet al. 1999); a conserved characterhas fewer changes,or "steps,"thanexpectedbasedon the numberof occurrencesin the terminaltaxa. Such tests areeasily modifiedto handleorderedor continuouscharacters(the Quantitative ConvergenceIndex;Ackerly & Donoghue 1998, Prinzing et al. 2001). An alternativemethod for continuoustraitsis based on regressions of traitdifferencesversusphylogeneticdistance(Legendreet al. 1994); a positive correlationindicates that traitsare conserved.The phylogenetic "neighborhood" over which trait conservatismis evident can be assessed with phylogenetic "autocorrelation"(Cheverudet al. 1985, Gittlemanet al. 1998, Bohning-Gaese & Oberrath1999). Owing to the nonparametric structure phylogenetic data, sigof nificancetesting often requiresrandomization methodsor null model simulations (Legendreet al. 1994, Lapointe& Garland2001). of Despite the proliferation such tests, few comparisonshavebeen conductedto evaluatetheirperformanceon common data sets. Morales (2000) conductedone suchcomparison, Ackerlyhasreanalyzed datawith severaladditional and his methods (D. Ackerly,unpublisheddata).Ackerly found thathierarchical ANOVA,the quantitative convergenceindex, andphylogeneticcorrelationof distancematrices give parallelresultsacrossdifferenttraits;however,therewas little correspondence between these methodsand phylogeneticautocorrelation eigenvectoranalyses. or For applicationsto communitydatasets, we favormethodsbased on phylogenetic distance (regression or autocorrelation) ratherthan parsimony-basedtrait mapping, to avoid the suggestionof inferringpatternsof historicaltraitevolutionfrom taxon sample.More work is needed to examine the stajust the community-based tistical power of these methodsand theirsensitivityto differentkinds of deviation from randompatterns.
Null Models for Community Phylogeniesand for CommunityAssembly

There has been extensive work on the generationof randomphylogenies (e.g., Raup et al. 1973, Losos & Adler 1995, Heard& Mooers 2000), but few studies have employed an explicitly ecological model (but see Maley 1998, Doebeli & Dieckmann2000). Hubbell(2001) has claimed thata realisticnull model for phylogenies must include informationon a region's biota (his "meta-community"), because the probabilityof a taxon'sextinctionis inverselyrelatedto its population size, and the sum of all populationsof all extant taxa is often limited (e.g., for canopy trees). The probabilityof extinctionis thereforedependenton the number of species in the region because increasingspecies richness will tend to increase
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ET WEBB AL. the numberof species with small populations.Hubbell's (2001) null models for phylogenies generatepatternsof hierarchicaldiversity (e.g., frequencydistributions of species per family) that fit observeddata well. Jansen & Mulder(1999) incorporated speciationinto a patch-dynamicmodel to simulatethe evolution of in an explicitly competitiveenvironment. lineages The simplest null models for communityphylogenies are generatedby subsamplingthe taxa in a largerarea,using existing phylogeniesfor the relationships among those taxa (e.g., Webb 2000). The large literatureon null models for the co-occurrence and patterns asassemblyof communitiesused to detectnonrandom is sembly rules (usually independentof phylogeny/taxonomy) relevanthere (e.g., Diamond 1975, Connor& Simberloff1979; reviewedby Gotelli & Graves 1996). of Null models also exist for the distribution ecological traitsexpectedin communities where competitionis important(Colwell & Winkler 1984, Leibold 1998, Stevens& Willig 2000) andfor the evolutionof traitsin lineages withouteffects of need to be combinedin future competition(e.g., Ackerly 2000); these approaches models.
Metricsof Community PhylogeneticStructure

Metrics that quantifythe distributionof taxa in a sample relative to a pool have been developedby Webb(2000). The net relatednessindex (NRI) is a standardized measureof the mean pairwisephylogeneticdistanceof taxa in a sample, relative to a phylogeny of an appropriate species pool, and quantifiesoverallclusteringof taxa on a tree (similarto Clarke& Warwick's1999 and von Euler & Svensson's 2001 metrics). It is calculated as -1 - ((mn(Xobs) - mnX(n))/sdX(n)), where Xobs is the phylogeneticdistancebetween two taxa (the sum of all interveningbranch is lengths) in the phylogeny of the pool, mn(Xobs) the mean of all possible pairs deviationexpected of n taxa, and mnX(n) and sdX(n) are the mean and standard for n taxa randomlydistributedon the phylogeny of the pool (found by multiple iteration;note thatthis formulationis slightly modifiedfrom Webb2000). Where continuousbranchlength estimates are not available,phylogeneticdistances can two be basedon the numberof nodes separating taxa(Farris1969, Gittleman& Kot measureof the phylogenetic 1990). Thenearesttaxonindex (NTI)is a standardized distanceto the nearesttaxon for each taxon in the sampleand quantifiesthe extent of terminalclustering,independentof deep level clustering.NTI is calculatedas
-1
.
is ((mn(Yobs)- mnY(n))/sdY(n)), where Yobs the phylogenetic distance to the
nearest taxon in the phylogeny of the pool; mn(Yobs), mnY(n), and sdY(n) are calculatedas for X. These metricssharemuchin commonwith those developedto assess the phylogenetic uniquenessof taxa in a conservationarea(e.g., Williamset al. 1991, Faith 1996, Crozier 1997, Nee & May 1997, Clarke& Warwick 1999, Sechrestet al. 2002). Both NRI andNTI increasewith increasingclusteringandbecome negative The with overdispersion. preciseresponseof NRI andNTI in communitiesformed attraction(Table 1) dependsupon the form the traitconservatism by phenotypic
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takes. Maximumconservatismin traits,at a deep level (leading to a high consisat tency index), yields both high NRI andNTI. Conservatism moreterminallevels in the phylogeny causes NTI to increasein significancerelativeto NRI. Both NRI and NTI depend on the particularspecies pool, and furtherstudy is requiredto determinewhen andhow these measurescan be comparedacrossdifferentstudies. A suitablenull model of communityassembly (see above) can be used to generate expectationsfor the distributionof relatednessindices with which the observed values can be compared.An alternativeapproachto assessing whetherthe taxa that co-occur in samples are more relatedthanexpected by chance is to correlate a metricof co-occurrencewith phylogeneticdistancefor all possible pairsof taxa (H. Steers,personalcommunication).
DIRECTIONS FOR FUTURE WORK

Beyond the directionsalreadytakenandreviewedin this paper,we have identified a numberof areasthatmight be profitableto pursue.
Dynamics of Community Phylogenetic Structure

The static patternsof communityphylogenetic structure describedabove (Webb 2000; H. Steers personalcommunication,J. Cavender-Bares personalcommunication) result from differentialmortalityof species that vary in phylogenetic relatednessand ecological characteristics. could Changesin phylogenetic structure also be observed directly over time in the many existing long-term datasets of communitycomposition.In age- and size-structured populations,comparingthe of communitystructure differentage- or size-classes at a single time can providea (limited)proxy for the directobservationof change over time (e.g., Webb& Peart 1999). Forexample,increasingsize classes of seedlings andtrees in small plots in Borneanrain forest shows a monotonicincreasein phylogeneticclustering(C.O. Webb,unpublisheddata).This pattern,at a single time, is consistentwith the cumulativemortalityof locally ill-suited species over time (if ecological suitability is phylogeneticallyconserved).
Using Phylogenetic Information in Models of Neighborhood Performance

Most models of the performance response(growth,survival)of focal individualsto neighborhooddensity classify neighborseitheras conspecifics or heterospecifics. This dichotomyhides a greatrangeof ecological similaritybetween species andan expected range of magnitudeof effect. Weightingthe interactionby a measureof of phylogeneticrelatednessshouldgreatlyimprovethe performance such models, if importantparametersof ecological similarityare phylogeneticallyconserved. For instance,if (a) negativeneighborhoodinteractionsaremediatedby pathogens or herbivores(e.g., Gilbertet al. 1994), (b) at least some of the pathogenspecies infect multiple host species, and (c) at least some of the polyphagouspathogens
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WEBBETAL. have a phylogeneticallyrestrictedset of host species (Futuyma& Mitter 1996, Farrell2001, Frenzel & Brandl 2001, Novotny et al. 2002), then the expected effect of neighbor density will be greater the more closely related it is to the focal individual. Analytical models of community stability,based on modified Lotka-Volteracompetition models with phylogenetically structuredinteraction coefficients;may also be possible andwouldbe expectedto predictthemaintenance of a phylogeneticallydiverse (or overdispersed) of species. set
ComparativeSurveysof Community PhylogeneticStructure

variationin communityphylogenetic structure across known graUnderstanding dients (e.g., moistureregime and species richness) may yield importantinsights into community organization(Thompson et al. 2001). Including gradientsthat integrate change in both resources and predation (e.g., Leibold 1996) will be reespecially revealing.The basic analysis of communityphylogenetic structure quires only plot-based samples and a species list (which can be convertedinto a communitysupertree),and can thus be rapidly conductedon preexistingdata. for Wherephylogeniescan be constructed fossils (e.g., Upchurch1995, Vermeij& turnoverof communitiescan be reconstructed Carlson2000) and a stratigraphic (e.g., Olszewski & Patzkowsky2001, Jackson& Overpeck2000), changein comcould be assessed over time. munityphylogenetic structure
PhylogeneticOrdinationand Classification
distancesthatreflect Basing ordinationand classificationmethodson intersample net phylogenetic dissimilarityratherthan Euclidean distance in N-dimensional species space offers a means to display the phylogeneticrelationsamong sampleplots. Such methods can reveal meaningful ecological relationshipshidden by standard,nonphylogeneticmethods:e.g., plots sharingmany genera should still clustereven if they sharenone of the same species.
Balanceof Community Phylogenies

Tree balance (the degree to which sister clades differ in their number of taxa) of providesanotherway to quantifythe complex branchingstructure community phylogenies(e.g., Heard& Mooers2000). Modelsrelatingthe phylogeneticdistriof butionof niche space amongtaxain a regionalpool andthe niche structure local communitiesshould generatepredictionsabout the balance of local community phylogenies.
CONCLUSIONS
We resist the temptationto declare that "phylogeneticcommunityecology" is a new field. Rather,we view phylogenetic informationas a "glue" that can stick ecological and evolutionarystudies together,where often they have slid past each
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other, their practitionersspeaking different languages. We want to emphasize, however, that despite its great utility, there is no simple, single way to apply in phylogeneticinformation communityecology, as is highlightedby the diversity of approachesreviewedhere. Phylogenies must also be used with full knowledge that of the assumptionsanduncertainties underliethem.Thereis a real dangerthat with the increasing ease of obtaining phylogenetic information,ecologists will forget thatphylogenies are hypothesesto be furthertested, and not the truth.This said, we genuinelybelieve thatno ecological studycan fail to benefitin some way of from an understanding the phylogenetic relationshipsof its taxa. Community and phylogeneticbiologists should continueto engage in a discussion ecologists that will surelyenrichand hopefully unite both disciplines. ACKNOWLEDGMENTS The synthesispresentedhere has grownout of discussionswith manypeople over the past fouryears,to whom we arevery grateful.We especially thankM. Ashton, P. Ashton, D. Baum, A. Blundell, J. Cavender-Bares,R. Chazdon, F. Cohen, J. Connell, R. Colwell, S. Davies, B. Enquist,T. Givnish, S. Heard,S. Hubbell, J. Losos, M. Martinez-Ramos, Peart,N. Pitman,R. Ree, M. Silman,P. Stevens, D. D. Wagner,M. Westoby,S. Zens, the Donoghue labs at Harvard Yale, a 1999 and seminar at Harvard,and an NCEAS working group on Life History graduate Strategiesof NeotropicalTrees. As the manuscriptwas being finalized, a meeting was held at NCEAS on "Phylogeniesand CommunityEcology" (organized by Donoghue & Webb), and we valued the opportunityto discuss many of the ideas in this paperwith those who attended.J. Losos, E. Palkovacs,andespecially B. Shaffer gave useful feedback on a draftof this manuscript.C.O.W.has been fundedby a MercerFellowshipof the ArnoldArboretum a Donnelley Fellowand of the Yale Institutefor BiosphericStudies. ship The Annual Reviewof Ecology and Systematics online at is http://ecolsys.annualreviews.org
LITERATURE CITED Ackerly DD. 1999. Comparative plant ecology and the role of phylogenetic information.In PhysiologicalPlant Ecology, ed. M Press,JD Scholes, MG Barker,pp. 391-413. Oxford, UK: Blackwell Sci. AckerlyDD. 2000. Taxonsampling,correlated evolution,and independentcontrasts.Evolution 54:1480-92 Ackerly DD, Donoghue M. 1998. Leaf size, sapling allometry,and Corer's rules: phylogeny and correlated evolution in maples (Acer).Am. Nat. 152:767-91 Ackerly DD, Dudley SA, Sultan SE, Schmitt J, Coleman JS, et al. 2000. The evolution of plant ecophysiological traits:recent advances and future directions. Bioscience 50:979-95 AshtonPS. 1988. Dipterocarp biology as a window to the understanding tropical forest of structure. Annu.Rev.Ecol. Syst. 19:347-70 Avise JC. 2000. Phylogeography:the History and Formationof Species. Cambridge,MA: Harvard Univ. Press Barker GM, Mayhill PC. 1999. Patterns of
498
WEBB ET AL. in Comparative Biology. Chicago: Univ. Chicago Press Brooks DR, McLennan DA. 1993. Historical ecology: examiningphylogeneticcomponents of communityevolution. See Ricklefs & Schluter1993a, pp. 267-80 Brooks DR, McLennanDA. 2002. TheNature of Diversity. Chicago:Univ. Chicago Press Brown JH. 1995. Macroecology. Chicago: Univ. ChicagoPress BrownJH,Zeng Z. 1989. Comparative population ecology of eleven species of rodentsin the Chihuahuan desert.Ecology 70:1507-25 Brown JM, McPeek MA, May ML. 2000. A phylogenetic perspective on habitat shifts and diversity in the North American Enallagma damselflies.Syst. Biol. 49:697-712 CadleJE, GreeneHW. 1993. Phylogeneticpatand terns,biogeography, the ecological structure of Neotropicalsnake assemblages. See Ricklefs & Schluter1993a, pp. 281-93 Cavender-Bares Holbrook NM. 2001. HyJ, draulic properties and freezing-induced xylem cavitationin evergreenanddeciduous oaks with contrastinghabitats. Plant, Cell Environ.24:1243-56 ChazdonRL, CareagaS, WebbCO, VargasO. 2002. Community and phylogenetic structure of reproductive traitsof woody species in wet tropicalforests.Ecol. Monogr.In press Chesson P. 2000. Mechanismsof maintenance of species diversity.Annu. Rev. Ecol. Syst. 31:343-66 Chesson PL, WarnerRR. 1981. Environmental variability promotescoexistencein lottery competitivesystems.Am. Nat. 117:923-43 CheverudJM, Dow MM, LeutneggerW. 1985. assessmentof phylogenetic The quantitative constraintsin comparativeanalyses: sexual in dimorphism body weight amongprimates. Evolution39:1335-51 ChownSL, GastonKJ.2000. Areas,cradlesand museums:the latitudinalgradientin species richness. TrendsEcol. Evol. 15:311-15 RM. 1999. The taxonomic ClarkeKR,Warwick distinctnessmeasureof biodiversity: weightleving of step lengths between hierarchical els. Mar.Ecol. Prog. Ser. 184:21-29
diversity and habitat relationships in terrestrialmollusc communitiesof the Pukeamaru ecological district, northeasternNew Zealand.J. Biogeogr.26:215-38 BarracloughTG, Hogan JE, Vogler AP. 1999. Testing whether ecological factors promote cladogenesis in a group of tiger beetles (Coleoptera: Cicindelidae). Proc. R. Soc. LondonSer.B 266:1061-67 TG, Barraclough Nee S. 2001. Phylogenticsand Ecol. Evol. 16:391-99 speciation.Trends TG, Vogler AP. 2000. DetectBarraclough ing the geographical pattern of speciation from species-level phylogenies. Am. Nat. 155:419-34 TG, VoglerAP, HarveyPH. 1998. Barraclough Revealing the factors that promote speciation. Philos. Trans. R. Soc. London Ser. B 353:241-49 Bininda-Emonds ORP,GittlemanJL, PurvisA. 1999. Buildinglargetreesby combiningphya logenetic information: completephylogeny of the extant Carnivora(Mammalia).Biol. Rev.74:143-75 Bininda-Emonds ORP,GittlemanJL, Steel M. 2002. The (super)tree of life: procedures, problems, and prospects. Annu. Rev. Ecol. Syst. 33:265-89 BlackburnTM, Gaston KJ. 2001. Local avian assemblagesas randomdrawsfromregional pools. Ecography24:50-58 Blondel J, Vigne J-D. 1993. Space, time and man as determinants diversityof birdsand of mammals in the Mediterranean region. See Ricklefs & Schluter1993a, pp. 135-46 R. B6hning-GaeseK, Oberrath 1999. Phylogenetic effects on morphological,life-history, behavioural and ecological traits of birds. Evol. Ecol. Res. 1:347-64 Bowers MA, Brown JH. 1982. Body size and coexistencein desertrodents:chanceor comEcology 63:391-400 munitystructure? Brandl R, Kristin A, Leisler B. 1994. Dietaryniche breadthin a local-communityof passerine birds, an analysis using phylogenetic contrasts.Oecologia 98:109-16 Brooks DR, McLennanDA. 1991. Phylogeny, Ecology,andBehavior:A ResearchProgram

Cody ML, Mooney HA. 1978. Convergence versus non-convergencein Mediterraneanclimate ecosystems. Annu. Rev. Ecol. Syst. 9:265-321 Colwell RK, WinklerDW. 1984. A null model for null models in biogeography. EcologiIn cal Communities: ConceptualIssues and the Evidence, ed. DR Strong,D Simberloff,LG Abele, AB Thistle, pp. 344-59. Princeton, NJ: PrincetonUniv. Press Comes HP, KadereitJW. 1998. The effects of Quaternaryclimatic changes on plant distribution and evolution. Trends Plant Sci. 3:432-38 ConnellJH.1980. Diversityandthecoevolution of competitors,or the ghost of competition past. Oikos 35:131-38 ConnorEF, SimberloffD. 1979. The assembly of species communities:chance or competition? Ecology 60:1132-40 CornellHV, LawtonJH. 1992. Species interactions, local andregionalprocesses,andlimits to the richness of ecological communities:a theoreticalperspective.J. Anim.Ecol. 61:112 CotgreaveP, HarveyPH. 1994. Phylogeny and the relationship betweenbody size andabundance in bird communities. Funct. Ecol. 8:219-28 Craig DA, Currie DC, Joy DA. 2001. Geographicalhistory of the central-western pacific blackfly subgenusInseliellum(Diptera: Simuliidae: Simulium) based on a reconstructedphylogeny of the species, hot-spot archipelagoes and hydrological considerations. J. Biogeogr.28:1101-27 CronquistA. 1988. The Evolutionand Classification of Flowering Plants. New York:NY Bot. Gard. CrozierRH. 1997. Preservingthe information content of species: genetic diversity, phylogeny, and conservationworth.Annu. Rev. Ecol. Syst. 28:243-68 CunninghamCW, Omland KE, Oakley TH. 1998. Reconstructing ancestral character states: a critical reappraisal.Trends Ecol. Evol. 13:361-66 Darwin C. 1859. The Origin of Species by
499
Means of Natural Selection. London: Murray DiamondJM. 1975. Assembly of species communities.In Ecology and Evolutionof Communities, ed. ML Cody, JM Diamond, pp. 342-444. Cambridge, MA: HarvardUniv. Press Dimichele WA,PhillipsTL. 1996. Clades,ecological amplitudes,and ecomorphs:phylogenetic effects and persistence of primitive plantcommunitiesin the Pennsylvanian-age tropicalwetlands.Palaeogeogr.Palaeoclim. Palaeoecol. 127:83-105 Dodd M, SilvertownJ, Chase M. 1999. Phylogenetic analysisof traitevolutionandspecies diversityvariationamong Angiospermfamilies. Evolution53:732-44 Doebeli M, DieckmannU. 2000. Evolutionary branchingand sympatricspeciation caused by differenttypes of ecological interactions. Am.Nat. 156:S77-101 Donoghue MJ, Bell CD, Li J. 2001. Phylogenetic patternsin NorthernHemisphereplant geography.Int. J. Plant Sci. 162:S41-52 Donoghue MJ, Ree RH. 2000. Homoplasyand developmentalconstraint:a model and example from plants.Am. Zool. 40:759-69 Donoghue MJ, Ree RH, Baum DA. 1998. Phylogeny andthe evolutionof flowersymmetry in the Asteridae.Trends Plant Sci. 3:311-17 EdwardsW, Westoby M. 2000. Families with highest proportionsof rare species are not consistent between floras. J. Biogeogr. 27: 733-40 RI, Eggleton P, Vane-Wright eds. 1994. Phylogenetics and Ecology. London:Academic Elton C. 1946. Competitionand the structure of ecological communities. J. Anim. Ecol. 15:54-68 Enquist BJ, Haskell JP, Tiffney BH. 2002. Generalpatternsof taxonomic structure and biomass division within extant and fossil woody plant communities.Nature.In press Faith DP. 1996. Conservation priorities and phylogenetic pattern. Conserv. Biol. 10: 1286-89 FarrellBD. 2001. Evolutionary assemblyof the milkweed fauna: cytochromeoxidase I and
500
WEBB ET AL. (India) using canonical correlationanalysis of herbarium records.Ecography.In press Gittleman JL, Anderson CG, Cates SE, Luh H-K, Smith JD. 1998. Detecting ecological patternsin phylogenies. In BiodiversityDynamics: Turnover Populations,Taxa,and of ed. Communities, ML McKinney,JA Drake, pp. 51-69. New York:ColumbiaUniv. Press statisGittlemanJL, Kot M. 1990. Adaptation: tics anda null model for estimatingphylogenetic effects. Syst. Zool. 39:227-41 Givnish T. 1998. Adaptiveradiationof plants molecon oceanic islands:classical patterns, ular data, new insights. In Evolution on Islands, ed. P Grant,pp. 281-304. Oxford, UK: OxfordUniv. Press GormanOT. 1992. Evolutionaryecology and historical ecology: assembly, structure,and organizationof streamfish communities.In Systematics,Historical Ecology, and North AmericanFresh WaterFishes, ed. RL Mayden, pp. 659-88. Stanford, CA: Stanford Univ. Press Gotelli NJ, Graves GR. 1996. Null Models in Inst. DC: Smithsonian Ecology. Washington, Press Gotelli NJ, TaylorCM. 1999. Testingmacroecological models with stream-fish assemblages. Evol. Ecol. Res. 1:847-58 GrandcolasP. 1993. The origin of biological diversity in a tropical cockroach lineage: a phylogenetic analysis of habitatchoice and biome occupancy.Acta Oecol. 14:259-70 GrandcolasP. 1998. Phylogeneticanalysis and the study of community structure. Oikos 82:397-400 Graves GR, Gotelli NJ. 1993. Assembly of avian mixed-species flocks in Amazonia. Proc. Natl. Acad. Sci. USA90:1388-91 Gregory RD. 1995. Phylogeny and relations among abundance,geographicalrange and body size of British breedingbirds. Philos. Trans.R. Soc. LondonSer.B 349:345-51 Grime JP. 1984. The ecology of species, families and communitiesof the contemporary Britishflora.New Phytol. 98:15-33 Hadly EA, MaurerBA. 2001. Spatialand temof poralpatterns species diversityin montane
the age of Tetraopes beetles. Mol. Phylogenet. Evol. 18:467-78 FarrellBD, DussourdDE, MitterC. 1991. Escalationof plant defense: Do latex and resin canals spur plant diversification?Am. Nat. 138:881-900 FarrisJS. 1969. A successive approximations weighting. Syst. Zool. approachto character 18:374-85 FelsensteinJ. 1985. Phylogenies and the comparativemethod.Am.Nat. 125:1-15 Fjeldsa J, Lovett JC. 1997. Geographicalpatterns of old and young species in African forestbiota:the significanceof specificmontaneareasas evolutionary centres.Biodivers. Conserv.6:325-46 ForstmeierW, Bourski OV, Leisler B. 2001. the Habitatchoice in Phylloscopuswarblers: role of morphology,phylogeny and competition. Oecologia 128:566-76 Fox BJ. 1999. The genesis and devlopmentof guild assemblyrules.InEcologicalAssembly Rules: Perspectives,Advances,Retreats,ed. E Weiher,P Keddy,pp. 23-57. Cambridge, UK: CambridgeUniv. Press Frenzel M, BrandlR. 2001. Hosts as habitats: insects befaunalsimilarityof phytophagous tween host plants. Ecol. Entomol. 26:594601 interFutuymaDJ, MitterC. 1996. Insect-plant actions:the evolutionof componentcommunities. Philos. Trans.R. Soc. London Ser. B 351:1361-66 diGalassiDMP.2001. Groundwatercopepods: versity patternsover ecological and evolutionaryscales. Hydrobiologia453:227-53 GardeziT, da Silva J. 1999. Diversity in relation to body size in mammals:a comparative study.Am.Nat. 153:110-23 of GentryAH. 1982. Patterns Neotropicalplant species diversity.Evol. Biol. 15:1-84 GilbertGS, HubbellSP,FosterRB. 1994. Deneffects of a canker sity and distance-to-adult disease of treesin a moist tropicalforest.Oecologia 98:100-8 C, Gimaret-Carpentier DrayS, PascalJ-P.2002. Broad-scalebiodiversitypatternanalyses of the endemic tree flora of the WesternGhats
AND COMMUNITYECOLOGY PHYLOGENIES mammal communities of western North America.Evol. Ecol. Res. 3:477-86 HarveyPH, Nee S. 1994. Comparingreal with expected patternsfrom molecular phylogenies. See Eggleton & Vane-Wright 1994, pp. 659-88 Harvey PH, Pagel MD. 1991. The Comparative Method in EvolutionaryBiology. Oxford, UK: OxfordUniv. Press Harvey PH, Rambaut A. 2000. Comparative analyses for adaptiveradiations.Philos. Trans.R. Soc. LondonSer.B 355:1599-605 Heard SB, Hauser DL. 1995. Key evolutionary innovationsand their ecological mechanisms. Hist. Biol. 10:151-73 Heard SB, Mooers AO. 2000. Phylogenetically patternedspeciation rates and extinction riskschangethe loss of evolutionary historyduringextinctions.Proc. R. Soc. London Ser.B 267:613-20 HerreraCM. 1992. Historicaleffects and sorting processes as explanations for contemporary ecological patterns: character syndromesin Mediterranean woody plants.Am. Nat. 140:421-46 Hubbell SP. 2001. The UnifiedNeutral Theory ofBiodiversityandBiogeography.Princeton, NJ: PrincetonUniv. Press Huelsenbeck JP, RonquistF, Nielsen R, Bollback JP. 2001. Bayesian inference of phylogeny and its impact on evolutionarybiology. Science 294:2310-14 Hunn CA, UpchurchP. 2001. The importance of time/space in diagnosing the causality of phylogenetic events: towardsa "chronobiogeographical" paradigm.Syst.Biol. 50:391407 Ibarra-Manriquez G, Martinez-Ramos M, Oyama K. 2001. Seedling functional types in a lowland rain forest in Mexico. Am. J. Bot. 88:1801-12 JablonskiD. 1987. Heritabilityat the species level: analysisof geographicrangesof cretaceous mollusks. Science 238:360-63 JablonskiD, Sepkoski JJ. 1996. Paleobiology, communityecology, and scales of ecological pattern.Ecology 77:1367-78 JacksonST, OverpeckJT. 2000. Responses of
501
plant populationsand communitiesto environmental Pachangesof thelate Quaternary. leobiol. 26:194-220 Jansen VAA, Mulder GSEE. 1999. Evolving biodiversity.Ecol. Lett. 2:379-86 Kelly CK. 1999. On the relationshipbetween function and phylogeneticrelatedness:environmentalseverityandcommunitystructure. XVIIntl. Botan. Congr.(Abstr.) F. Kelly CK, Woodward 1996. Ecological correlates of plant range size: taxonomies and phylogenies in the study of plant commonness andrarityin greatbritain.Philos. Trans. R. Soc. LondonSer.B 351:1261-69 Lapointe F-J, Cucumel G. 1997. The average consensus procedure: combination of weighted trees containingidentical or overlapping sets of taxa. Syst. Biol. 46:306-12 Lapointe F-J, GarlandT. 2001. A generalized model for the analysis of crosspermutation species data.J. Classif. 18:109-27 Latham RE, Ricklefs RE. 1993. Continental comparisonsof temperate-zonetree species diversity.See Ricklefs & Schluter1993a,pp. 294-314 Lechowicz MJ. 1984. Why do temperate deciduous trees leaf out at differenttimes? Adaptations and ecology of forest communities. Am.Nat. 124:821-42 P. LegendreP,LapointeF, Casgrain 1994. Modeling brain evolution from behavior:a permutational regression approach. Evolution 48:1487-99 Leibold MA. 1996. A graphicalmodel of keystone predatorsin food webs: trophicregulation of abundance,incidence and diversity patternsin communities.Am. Nat. 147:784812 Leibold MA. 1998. Similarity and local coexistence of species in regionalbiotas. Evol. Ecol. 12:95-110 Lewis PO. 2001. Phylogenetic systematics turns over a new leaf. TrendsEcol. Evol. 18:30-37 LindemanPV. 2000. Resourceuse of five sympatric turtle species: effects of competition, phylogeny, and morphology. Can. J. Zool. 78:992-1008
502
WEBB ET AL. Maley CC. 1998. Theevolutionof biodiversity: a simulation approach. PhD thesis. Cambridge,MA: MIT Mayden RL. 1987. Historical ecology and North Americanhighlandfishes: a research program in community ecology. In Communityand EvolutionaryEcology of North AmericanStreamFishes, ed. WJ Matthews, DC Heins, pp. 210-22. Norman,OK: Univ. Okla. Press Mazer SJ, WheelwrightNT. 1993. Fruit size and shape: allometryat differenttaxonomic levels in bird-dispersedplants. Evol. Ecol. 7:556-75 McCallum DA, Gill FB, Gaunt SLL. 2001. Community assembly patterns of parids along an elevational gradient in western China.WilsonBull. 113:53-64 McPeekMA. 1996. Linkinglocal species interactionsto ratesof speciationin communities. Ecology 77:1355-66 McPeek MA, Brown JM. 2000. Building a regional species pool: diversificationof the Enallagma damselflies in easternNorth Americanwaters.Ecology 81:904-20 McPeek MA, Miller TE. 1996. Evolutionary biology and community ecology. Ecology 77:1319-20 Miles DB, Dunham AE. 1993. Historical perspectives in ecology and evolutionary biology-the use of phylogenetic comparative analyses.Annu.Rev.Ecol. Syst. 25:587619 Miller JS, Wenzel JW. 1995. Ecological characters and phylogeny. Annu. Rev. Entomol. 40:389-415 Morales E. 2000. Estimatingphylogenetic inertiain Tithonia(Asteraceae):a comparative Evolution54:475-84 approach. MorandS, Guegan JF. 2000. Distributionand abundanceof parasite nematodes: ecological specialisation,phylogeneticconstraintor simply epidemiology?Oikos 88:563-73 Moreau RE. 1948. Ecological isolation in a J. richtropicalavifauna. Anim.Ecol. 17:11326 MurrayBR, Westoby M. 2000. Propertiesof curves: species in the tail of rank-abundance
Lord J, Westoby M, LeishmanM. 1995. Seed size and phylogeny in six temperatefloras: constraints,niche conservatism,and adaptation. Am.Nat. 146:349-64 Losos JB. 1992. The evolution of convergent Anolis communities. structurein Caribbean Syst. Biol. 41:403-20 Losos JB. 1995. Community evolution in GreaterAntillean Anolis lizards: phylogenetic patternsandexperimental tests. Philos. Trans.R. Soc. LondonSer.B 349:69-75 Losos JB. 1996. Phylogenetic perspectiveson communityecology. Ecology 77:1344-54 Losos JB, Adler FR. 1995. Stumpedby trees:a generalizednull model for patternsof organismal diversity.Am.Nat. 145:329-42 Losos JB, JackmanTR, LarsonA, de Queiroz K, Rogriguez-Schettino L. 1998. Contingency and determinismin replicatedadaptive radiations of island lizards. Science 279:2115-18 Losos JB, Schluter D. 2000. Analysis of an evolutionary species-area relationship.Nature408:847-50 Losos JB, Warheit KI, Schoener TW. 1997. Adaptive differentiation following experimentalisland colonizationin Anolis lizards. Nature 387:70-73 LynchJD. 1989. The gaugeof speciation:on the frequencyof modes of speciation.In Speciation and Its Consequences,ed. D Otte, JA Endler, pp. 527-53. Sunderland,MA: Sinauer MacArthurRH. 1960. On the relative abundance of species. Am.Nat. 94:25-36 MacArthur RH, Levins R. 1967. The limiting similarity, convergenceanddivergenceof coexisting species. Am.Nat. 101:377-85 MacArthurRH, Wilson EO. 1967. The Theory of Island Biogeography.Princeton,NJ: PrincetonUniv. Press Maddison WP, Slatkin M. 1991. Null models for the number of evolutionary steps in a characteron a phylogenetic tree. Evolution 45:1184-97 MagallonS, SandersonMJ. 2001. Absolute diversification ratesin angiosperm clades.Evolution 55:1762-80
PHYLOGENIES AND COMMUNITYECOLOGY the potentialfor increasein abundance. Evol. Ecol. Res. 2:583-92 Nee S, May RM. 1997. Extinction and loss of evolutionaryhistory.Science 278:692-95 Nee S, Read AF, GreenwoodJJD,HarveyPH. 1991. Phylogeny and the relationship between abundanceand body size in British birds.Nature 351:312-13 Nel A, Nel J, Masselot G, ThomasA. 1998. An investigationinto the applicationof the Wagnerparsimonymethodin synecology.Biol. J. Linn. Soc. 65:165-89 Novotny V, Basset Y, Miller SE, Weiblen G, BremerB, et al. 2002. Low host specificity of herbivorousinsects in a tropical forest. Nature416:841-44 Olszewski TD, Patzkowsky ME. 2001. Evaluating taxonomic turnover:PennsylvanianPermian brachiopods and bivalves of the NorthAmericanmidcontinent. Paleobiology 27:646-68 Peat HJ, Fitter AH. 1994. Comparativeanalyses of ecological characteristicsof British angiosperms.Biol. Rev. 69:95-115 Peterson AT, Soberon J, Sanchez-CorderoV. 1999. Conservatismof ecological niches in evolutionarytime. Science 285:1265-67 Poulin R. 1999. Speciation and diversification of parasitelineages: an analysis of conEvol. generic parasitespecies in vertebrates. Ecol. 13:455-67 Price T, Lovette IJ, BerminghamE, Gibbs HL, RichmanAD. 2000. Theimprint historyon of communitiesof North American and Asian warblers.Am. Nat. 156:354-67 PrinzingA, DurkaW, Klotz S, BrandlR. 2001. The niche of higherplants:evidencefor phyProc. R. Soc. London logenetic conservatism. Ser.B 268:2383-89 Qian H, Ricklefs RE. 1999. A comparisonof the taxonomic richness of vascular plants in China and the United States. Am. Nat. 154:160-81 Raup DM, Gould SJ, Schopf TJM, Simberloff DS. 1973. Stochastic models of phylogeny and the evolution of diversity. J. Geol. 81:525-42 Richardson Pennington Pennington TD, JE, RT,
503
HollingsworthPM. 2001. Rapid diversification of a species-rich genus of Neotropical rain forest trees. Science 293:2242-45 RichmanAD. 1996. Ecological diversification and community structurein the Old World leaf warblers(genusPhylloscopus): a phylogenetic perspective. Evolution 50:246170 RichmanAD, Price T. 1992. Evolutionof ecological differencesin the Old Worldleaf warblers. Nature 355:817-21 Ricklefs RE. 1987. Communitydiversity:relative roles of local and regional processes. Science 235:167-71 Ricklefs RE. 2002. Phylogenetic perspectives on patterns regionalandlocal species richof ness. In TropicalRainforests:Past and Future,ed. C Moritz,E Bermingham.In press Ricklefs RE, BerminghamE. 2001. Nonequilibriumdiversitydynamicsof the LesserAntillean avifauna.Science 294:1522-24 Ricklefs RE, LathamRE. 1992. Intercontinental correlationof geographicalranges suggests stasis in ecological traitsof relict genera of temperateperennialherbs. Am. Nat. 139:1305-21 Ricklefs RE, SchluterD, eds. 1993a. Species Diversity in Ecological Communities:Historical and Geographical Perspectives. Chicago:Univ. Chicago Press Ricklefs RE, SchluterD. 1993b. Species diversity: regional and historical influences. See Ricklefs & Schluter1993a, pp. 350-63 SalaminN, HodkinsonTR, SavolainenV. 2002. an Buildingsupertrees: empiricalassessment using the grass family (Poaceae).Syst. Biol. 51:136-50 Sanderson MJ, Donoghue MJ. 1996. Reconstructing shifts in diversification rates on Ecol. Evol. 11:15phylogenetictrees.Trends 20 SandersonMJ, PurvisA, Henze C. 1998. Phylogenetic supertrees: assemblingthe trees of life. TrendsEcol. Evol. 13:105-9 Sanmartin I, Enghof H, Ronquist F. 2001. Patternsof animal dispersal,vicariance and in diversification the Holarctic.Biol. J. Linn. Soc. 73:345-90
504
WEBB ET AL. in tropical rain forest in Guyana. Ecology 82:3197-212 ThompsonJN, ReichmanOJ, Morin PJ, Polis of GA, PowerME, et al. 2001. Frontiers ecology. Bioscience 51:15-24 Tilman D. 1994. Competition and biodiverhabitats.Ecology sity in spatiallystructured 75:2-16 ToftsR, SilvertownJ. 2000. A phylogeneticapproachto communityassembly from a local species pool. Proc. R. Soc. London Ser. B 267:363-69 Tokeshi M. 1991. Faunal assembly in chironomids(Diptera):generic associationand spread.Biol. J. Linn. Soc. 44:353-67 UpchurchP. 1995. The evolutionaryhistoryof sauropoddinosaurs.Philos. Trans.R. Soc. LondonSer.B 349:365-90 VerduM, Barron-SevillaJA, Valiente-Banuet A, Flores-HernandezN, Garcia-Fayos P. 2002. Mexican plant phenology: Is it similar to Mediterranean communities?Bot. J. Linn. Soc. 138:297-303 VermeijGJ,CarlsonSJ. 2000. The muricidgastropodsubfamilyRapaninae: phylogeny and ecological history.Paleobiology 26:19-46 VittLJ, Zani PA. 1996. Organization a taxoof nomicallydiverselizardassemblagein Amazonian Ecuador.Can. J. Zool. 74:1313-35 von EulerF, Svensson S. 2001. Taxonomicdistinctness and species richness as measures of functional structurein bird assemblages. Oecologia 129:304-11 HE, Wanntorp Brooks DR, Nilsson T, Nylin S, Ronquist F. 1990. Phylogenetic approaches in ecology. Oikos 57:119-32 E. Warming 1909. Oecologyof Plants. London: OxfordUniv. Press Webb CO. 2000. Exploring the phylogenetic structureof ecological communities:an exampleforrainforesttrees.Am.Nat. 156:14555
Sasal P, Niquil N, BartoliP. 1999. Community structure digeneanparasitesof sparidand of labridfishes of the Mediterranean a new sea: 119:635-48 Parasitology approach. Schluter D. 2000a. Ecological characterdisplacement in adaptive radiation.Am. Nat. 156:S4-16 RaSchluterD. 2000b.TheEcology ofAdaptive diation. Oxford,UK: OxfordUniv. Press Schluter D. 2001. Ecology and the origin of species. TrendsEcol. Evol. 16:372-80 Schluter D, Ricklefs RE. 1993. Convergence and the regional component of species diversity. See Ricklefs & Schluter 1993a, pp. 230-41 SchwarzMW, SimberloffD. 2001. Taxon size predicts rates of rarity in vascular plants. Ecol. Lett.4:464-69 Sechrest W, Brooks TM, da Fonseca GAB, Konstant WR, Mittermeier RA. 2002. Hotspots and the conservationof evolutionary history.Proc. Natl. Acad. Sci. USA 99: 2067-71 Semple C, Steel M. 2000. A supertreemethod for rooted trees. Discrete Appl. Math. 105: 147-58 SilvertownJ, Dodd ME, Gowing DJG. 2001. of Phylogenyandtheniche structure meadow plantcommunities.J. Ecol. 89:428-35 SilvertownJ, Dodd ME, Gowing DJG, MountfordJO. 1999. Hydrologicallydefinedniches reveala basis for species richnessin communites. Nature400:61-63 Simberloff DS. 1970. Taxonomic diversity of island biotas. Evolution24:23-47 Stevens RD, Willig MR. 2000. Community and Oikos abundance, morphology. structure, 88:48-56 SturmbauerC. 1998. Explosive speciation in cichlid fishes of the African great lakes: a J. dynamicmodel of adaptiveradiation. Fish Biol. 53:18-36 Taberlet P, Fumagalli L, Wust-Saucy A, Cosson J. 1998. Comparativephylogeography and postglacial colonization routes in Europe.Mol. Ecol. 7:453-64 ter Steege H, Hammond DS. 2001. Character convergence, diversity, and disturbance
Webb CO, Donoghue MJ. 2002. Phylomatic: A Database for Applied Phylogenetics. http://www.phylodiversity.net/phylomatic WebbCO,PeartDR. 1999. Seedlingdensitydependence promotescoexistence of Bomean rain forest trees. Ecology 80:2006-17
AND COMMUNITYECOLOGY PHYLOGENIES WebbCO, PeartDR. 2000. Habitatassociations of trees and seedlings in a Bomean rain forest. J. Ecol. 88:464-78 WebbCO, PitmanNC. 2002. Phylogeneticbalance and ecological evenness. Syst. Biol. In press WestobyM, LeishmanM, LordJ. 1995. Onmisthe J. interpreting "phylogeneticcorrection." Ecol. 83:531-34 Williams CB. 1964. Patternsin the Balance of Nature.New York: Academic Williams PH, HumphriesCJ, Vane-Wright RI. 1991. Measuringbiodiversity:taxonomicrelatedness for conservationpriorities.Austr. Syst. Bot. 4:665-79 Williams-Linera 1997. Phenology of decidG. uous andbroadleaved-evergreen species tree in a Mexican tropicallower montaneforest. Glob. Ecol. Biogeogr.Lett. 6:115-27
505
Willis JC. 1922.Area andAge: A Studyof Geoand graphicalDistribution Originin Species. Cambridge,UK: CambridgeUniv. Press Wing SL, Hickey LJ, Swisher CC. 1993. Late Cretaceousvegetation.Nature 363:342-44 WinstonMR. 1995. Cooccurrenceof morphologically similarspecies of streamfishes.Am. Nat. 145:527-45 Wisheu IC. 1998. How organisms partition habitats: different types of community organization can produce identical patterns. Oikos 83:246-58 ZimmermanBL, SimberloffD. 1996. An historical interpretation habitatuse by frogs of in a CentralAmazonianforest. J. Biogeogr. 23:27-46 Zinc RM. 1996. Comparative phylogeography in NorthAmericanbirds.Evolution50:30817

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Phylogenies and Community Ecology Author(s): Campbell O. Webb, David D. Ackerly, Mark A. McPeek and Michael J.

AND COMMUNITY PHYLOGENIES ECOLOGY

COMMUNITY STRUCTURE AND COEXISTENCE

PHYLOGENIES COMMUNITY AND ECOLOGY

Community Niche Structure

The PhylogeneticStructureof CommunityAssemblages

ECOLOGY AND PHYLOGENIES COMMUNITY

(iv) Community Assembly:

Fundamental Niche Y Fundamental NicheZ ^ ECOLOGY

AND PHYLOGENIES COMMUNITY ECOLOGY

The PhylogeneticBasis of CommunityNiche Structure

AND PHYLOGENIES COMMUNITY ECOLOGY

BiogeographicHistory,CharacterEvolution, and CommunityAssembly

AND COMMUNITYECOLOGY PHYLOGENIES

AND PHYLOGENIES COMMUNITY ECOLOGY

AND ECOLOGY PHYLOGENIES COMMUNITY

EMERGENT PROPERTIES COMMUNITIES OF

AND PHYLOGENIES COMMUNITY ECOLOGY

ADVANCES METHODOLOGICAL Community Phylogenies

PHYLOGENIES COMMUNITY AND ECOLOGY

Null Models for Community Phylogeniesand for CommunityAssembly

Metricsof Community PhylogeneticStructure

is ((mn(Yobs)- mnY(n))/sdY(n)), where Yobs the phylogenetic distance to the

AND ECOLOGY PHYLOGENIES COMMUNITY

DIRECTIONS FOR FUTURE WORK

Dynamics of Community Phylogenetic Structure

Using Phylogenetic Information in Models of Neighborhood Performance

ComparativeSurveysof Community PhylogeneticStructure

Balanceof Community Phylogenies

AND PHYLOGENIES COMMUNITY ECOLOGY

PHYLOGENIES COMMUNITY AND ECOLOGY

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